studies on the stoneflies of georgia with...

20
Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03 http://zoobank.org/ urn:lsid:zoobank.org:pub:F7668389-D0A8-4697-938F-C8D1DF8D129A Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 30 STUDIES ON THE STONEFLIES OF GEORGIA WITH THE DESCRIPTION OF A NEW SPECIES OF SOYEDINA RICKER, NEW STATE RECORDS AND AN ANNOTATED CHECKLIST Chris J. Verdone 1 , Boris C. Kondratieff 1 , R. Edward DeWalt 2 , and Eric J. South 3 1 Department of Bioagricultural Sciences and Pest Management, Colorado State University, Fort Collins, Colorado, 80523, U.S.A. E-mail: [email protected] E-mail: [email protected] 2 University of Illinois, Prairie Research Institute, Illinois Natural History Survey, 1816 S Oak St., Champaign, Illinois, 61820, U.S.A. E-mail: [email protected] 3 Department of Entomology, University of Illinois at Urbana-Champaign, 320 Morrill Hall, 505 S. Goodwin Ave., Urbana, Illinois, 61801, U.S.A. E-mail: [email protected] ABSTRACT A new species of Soyedina Ricker is described, in addition to the listing of 24 new state records and the inclusion of an annotated checklist of species presently known from Georgia. Keywords: Plecoptera, Nemouridae, Soyedina amicalola Verdone & Kondratieff, 2017, Georgia, checklist INTRODUCTION Georgia stoneflies have received relatively little attention from entomologists in recent years. Although 14 valid species have their type localities in Georgia (Table 1, DeWalt et al. 2017), no systematic review of the stonefly fauna of the state has been published. Presently, only 75 species of stoneflies have been reported from Georgia (DeWalt et al. 2017). By comparison, 107 species have been reported from adjacent Alabama (Grubbs 2011), an area of comparable size. Additions and modifications to the Georgia stonefly fauna have typically resulted from occasional taxonomic treatments, i.e., Taeniopteryx (Ricker & Ross 1968), Allocapnia (Ross & Ricker 1971), Acroneuria (Stark & Gaufin 1976), Chloroperlinae (Surdick 1985), and Isoperla (Szczytko & Kondratieff 2015). Previously, the greatest contribution to the stonefly fauna of Georgia resulted from Ricker (1952), in which 17 new state records were reported. The purpose of this paper is to present the description of a new species of Soyedina Ricker, report new state records and provide a preliminary checklist of the stoneflies of Georgia.

Upload: others

Post on 16-Jun-2020

1 views

Category:

Documents


0 download

TRANSCRIPT

Page 1: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the

description of a new species of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49.

https://doi.org/10.25031/2017/13.03

http://zoobank.org/ urn:lsid:zoobank.org:pub:F7668389-D0A8-4697-938F-C8D1DF8D129A

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 30

STUDIES ON THE STONEFLIES OF GEORGIA WITH THE DESCRIPTION OF

A NEW SPECIES OF SOYEDINA RICKER, NEW STATE RECORDS AND

AN ANNOTATED CHECKLIST

Chris J. Verdone1, Boris C. Kondratieff1, R. Edward DeWalt2, and Eric J. South3

1 Department of Bioagricultural Sciences and Pest Management, Colorado State University,

Fort Collins, Colorado, 80523, U.S.A.

E-mail: [email protected]

E-mail: [email protected]

2 University of Illinois, Prairie Research Institute, Illinois Natural History Survey, 1816 S Oak St.,

Champaign, Illinois, 61820, U.S.A.

E-mail: [email protected]

3 Department of Entomology, University of Illinois at Urbana-Champaign, 320 Morrill Hall, 505

S. Goodwin Ave., Urbana, Illinois, 61801, U.S.A.

E-mail: [email protected]

ABSTRACT

A new species of Soyedina Ricker is described, in addition to the listing of 24 new state records and the

inclusion of an annotated checklist of species presently known from Georgia.

Keywords: Plecoptera, Nemouridae, Soyedina amicalola Verdone & Kondratieff, 2017, Georgia, checklist

INTRODUCTION

Georgia stoneflies have received relatively little

attention from entomologists in recent years.

Although 14 valid species have their type localities

in Georgia (Table 1, DeWalt et al. 2017), no

systematic review of the stonefly fauna of the state

has been published. Presently, only 75 species of

stoneflies have been reported from Georgia (DeWalt

et al. 2017). By comparison, 107 species have been

reported from adjacent Alabama (Grubbs 2011), an

area of comparable size.

Additions and modifications to the Georgia

stonefly fauna have typically resulted from

occasional taxonomic treatments, i.e., Taeniopteryx

(Ricker & Ross 1968), Allocapnia (Ross & Ricker

1971), Acroneuria (Stark & Gaufin 1976),

Chloroperlinae (Surdick 1985), and Isoperla

(Szczytko & Kondratieff 2015). Previously, the

greatest contribution to the stonefly fauna of

Georgia resulted from Ricker (1952), in which 17

new state records were reported. The purpose of

this paper is to present the description of a new

species of Soyedina Ricker, report new state records

and provide a preliminary checklist of the stoneflies

of Georgia.

Page 2: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 31

Table 1. Species with type localities in Georgia.

Family Species Type

Chloroperlidae? Chloroperla cydippe Newman, 1839 Nomen dubium

Leuctridae Leuctra biloba Claassen, 1923 Holotype

Leuctridae Leuctra moha Ricker, 1952 Holotype

Peltoperlidae Tallaperla anna (Needham & Smith, 1916) Syntype

Peltoperlidae Tallaperla cornelia (Needham & Smith, 1916) Holotype

Peltoperlidae Tallaperla laurie (Ricker, 1952) Holotype

Peltoperlidae Viehoperla ada (Needham & Smith, 1916) Holotype

Perlidae Acroneuria petersi Stark & Gaufin, 1976 Holotype

Perlidae Beloneuria georgiana (Banks, 1914) Holotype

Perlidae Neoperla clymene (Newman, 1839) Holotype

Perlidae Perlinella drymo (Newman, 1839) Holotype

Perlidae Perlinella ephyre (Newman, 1839) Holotype

Perlodidae Clioperla clio (Newman, 1839) Holotype

Perlodidae Isoperla chickamauga Szczytko & Kondratieff, 2015 Holotype

Perlodidae Remenus duffieldi Nelson & Kondratieff, 1995 Holotype

METHODS

Adult stoneflies were collected in March and

May 2016 and February and May 2017 at locations

in the five physiographic provinces of Georgia

(Appalachian Plateau, Blue Ridge, Coastal Plain,

Piedmont Plateau, and Ridge and Valley) (Fig. 1,

Appendix A). Adult stoneflies were collected using

a beating sheet (BioQuip, Rancho Dominguez,

California, catalog # 2840C) or handpicked from

substrates using forceps. Specimens were preserved

in 80% or 90% ethanol. Locality data were recorded

directly using either Topo Maps version 1.16 for I-

Phone or a Garmin eTrex 20. Locality data for some

records were georeferenced using ACME Mapper

2.1 (https://mapper.acme.com) or GEOLocate v. 3.22

(http://www.museum.tulane.edu/geolocate).

Additional material were examined from the C. P.

Gillette Museum of Arthropod Diversity, Colorado

State University, Fort Collins, Colorado (CSUC),

Clemson University Arthropod Collection,

Clemson, South Carolina (CUAC), the Illinois

Natural History Survey, Champaign, Illinois

(INHS) and Western Kentucky University, Bowling

Green, Kentucky (WKUC). Specimens were

inspected under a dissecting stereomicroscope.

Color images were captured following the methods

of Verdone and Kondratieff (2016). Image

adjustments and measurements were made using

Adobe Photoshop CS6 Extended.

RESULTS

A total of 1,776 adult stoneflies were examined

in the study. The description of a new species of

Soyedina, 24 new state records and an annotated

checklist are presented below. Data for all

specimens examined can be downloaded from a

comma separated values file.

Soyedina amicalola Verdone & Kondratieff, sp. n.

http://lsid.speciesfile.org/urn:lsid:Plecoptera.speciesfil

e.org:TaxonName:499523

(Figs. 2-8)

Material examined. Holotype ♂, Georgia, Dawson

Co., Little Amicalola Creek, Amicalola Falls State

Park, N 34.57233, W 84.24107, 9 February 2017, C.

Verdone, B. Kondratieff, (USNM); Paratypes: Same

Page 3: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 32

Fig. 1. Map of the physiographic provinces of Georgia and 2016-2017 collecting locations.

data as holotype, ♂ (CSUC) (SCAN,

CSUC_ENT0062195); Lumpkin Co., Frogtown

Creek, Hwy 19, Desoto Falls Rec. Area, N 34.70649,

W 83.91634, 10 February 2017, C. Verdone, B.

Kondratieff, ♂ (CSUC) (SCAN, CSU_ENT0062194).

Distribution. USA – GA

Etymology. The name amicalola is a Cherokee word

meaning “tumbling waters” and is a reference to the

nearby waterfalls where this species was collected.

We propose Georgia Forest Fly as the common

name for this new species.

1

Page 4: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 33

Fig. 2. Soyedina amicalola sp. n. male dorsal habitus.

Male. Macropterous. Length of forewings 7.8–7.9

mm (n = 3). Length of body 6.4–6.6 mm (n = 3).

General body color brown (Fig. 2). Head dark

brown; U-shaped carina between ocelli. Ocelli

reddish. Legs light brown. Wings fumose. Venation

typical for genus with A1 and A2 united near wing

margin. Gills absent. Cerci membranous and one

segmented with a sclerotized nipple-like process on

apex. Hypoproct well developed, base covering

medial ⅓ of ninth sternum (Figs. 3, 6); apical half

articulated, bent dorsad; small lateral projections

covered in long setae produced at bend (Figs. 3, 5, 6,

8); apex lightly sclerotized, cylindrical. Vesicle long

and thin, 3.5X as long as wide (n = 3); narrow at base;

widest at basal ¼; tapering to a narrowly rounded

tip (Figs. 3, 6). Paraprocts with two lobes; inner lobes

inconspicuous, small, narrow, flattened and lightly

sclerotized. Outer lobes large, heavily sclerotized,

narrowed at midlength; apical portion expanded

posteriorly, mushroom-shaped in profile (Fig. 5, 8).

Apical half of proximal side of outer lobes covered

in sensilla, giving the appearance of a serrated edge

on apical margin (Figs. 3–5, 8). Anterior apex

broadly pointed and oriented slightly ventrad.

Apex bisected by a thin sclerotized band (Figs. 5, 8).

Epiproct typical for genus, asymmetrical with right

half larger than left (Figs. 3, 4, 6, 7). Epiproct 2.5X as

long as wide (n = 3), when measured from base of

ventral sclerite to epiproct apex and widest width.

Lateral arms short, extending from the base of

ventral sclerites to the base of the sclerotized portion

of the dorsal sclerites (Fig. 7). Ventral sclerites

elongate, width subequal, bearing small teeth on

outer margin, tips rounded (Figs. 4, 7). Basal

sclerites rectangular (Fig. 7). Basal portion of dorsal

sclerite darkly sclerotized, triangular shaped,

encircling basal cushion. Apical portion of dorsal

sclerite lightly sclerotized; open throughout the

apical ⅔, which exposes the inner sclerotized

structure.

Female. Unknown.

Diagnosis. Soyedina amicalola is most similar to S.

kondratieffi Baumann & Grubbs, 1996, S. merritti

Baumann & Grubbs, 1996 and S. washingtoni

(Claassen, 1923) which also possess pointed

paraproct apices, but can be separated by other

details of the paraprocts. Soyedina amicalola has

paraprocts that are mushroom-shaped with sensilla

covering the apical half of the proximal side, giving

the margin a serrated appearance. Whereas, S.

kondratieffi, S. merritti and S. washingtoni do not

2

Page 5: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 34

Figs. 3-5. Soyedina amicalola sp. n. 3. Male vesicle, ventral. 4. Male terminalia, dorsal. 5. Male terminalia,

lateral.

exhibit the mushroom-shape and serrated

appearance. In addition, S. amicalola is distinguished

from S. kondratieffi by details of the epiproct. The

epiproct of S. amicalola is asymmetrical, in contrast

to the nearly symmetrical epiproct of S. kondratieffi.

Ongoing work by Scott Grubbs and Richard

Baumann has found that details of epiprocts possess

characters useful in grouping eastern Nearctic

species of Soyedina (S. Grubbs, personal

communication). Based on these characters,

3 4

5

Page 6: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 35

Figs. 6-8. Soyedina amicalola sp. n. 6. Male vesicle, ventral. 7. Male terminalia, dorsal. 8. Male paraproct,

lateral. BS = basal sclerite, DS = dorsal sclerite, HP = hypoproct, ITP = inner tube-shaped process, LA = lateral

arm, OL = outer paraproct lobe, VC = vesicle, VS = ventral sclerite.

S. amicalola is tentatively grouped with S.

washingtoni, S. carolinensis (Claassen, 1923), and S.

merritti, all of which possess relatively straight

ventral sclerites and an inner tube-shaped process

that is flared apically with the opening mostly

centered, not strongly oriented to the left (Figs. 7, 8).

Soyedina amicalola is easily distinguished from S.

carolinensis which lacks pointed paraproct apices

and is truncate posteriorly.

Additional material examined: Soyedina

carolinensis: North Carolina, Buncombe Co., Blue

Ridge Parkway, MP 360.3, BLRI, 2 May 2007, J.

Robinson, ♂, ♀ (CSUC); Haywood Co., ATBI Plot,

Cataloochee, GSMNP, 27 March–10 April 2002, P.E.

Super, 2♂, ♀ (CSUC); ATBI Plot, Purchase Knob,

GSMNP, 27 March–10 April 2002, P.E. Super, J.

Lowe, ♂ (CSUC); Big Creek, GSMNP boundary,

11March 2008, C.R. Parker, ♂ (CSUC); Macon Co.,

stream at Wayah Crest Campground, Nantahala

National Forest, 5 March 1991, R. Baumann, S.

6 7

8

Page 7: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 36

Clark, 6♂, 2♀ (CSUC). Soyedina kondratieffi: North

Carolina, Macon Co., Upper Ball Creek, Coweeta

Hydro Lab, (Malaise) 20 March–7 April 1984, A.D.

Huryn, ♂ (CSUC, paratype); Swain Co., Andrews

Bald, ATBI plot, GSMNP, (Malaise), 16 March–24

April 2001, I.C. Stocks, J. Breeden, 4♂, 4♀ (CSUC);

Tennessee, Sevier Co., Rocky Spur of Roaring Fork,

GSMNP, 25 March 1999, B. Kondratieff, ♂ (CSUC);

Sams Creek, Stn 7, third trail crossing, ca. 100 m

below mouth of Chuck Hollow, GSMNP, 1 February

1997, D. Etnier, ♂ (CSUC); Mannis Branch, ca. 12 mi.

Little River, GSMNP, 25 February 1998, B. Hart, ♂

(CSUC). Soyedina merritti: Pennsylvania,

Westmoreland Co., Maul Spring, Powdermill

Nature Reserve, 19 March 1975, R.W. Baumann, O.S.

Flint Jr., J.L. Sykora ♂ (USNM, holotype). Soyedina

sp.: Georgia, Rabun Co., Greasy Creek, North

Germany Rd., N 34.9146, W 83.41972, 10 February

2017, C. Verdone, B. Kondratieff, ♀ (CSUC).

Soyedina washingtoni: New York, Delaware Co.,

spring-fed trib. to Emory Brook, Rt. 28

Fleischmanns, N 42.1511, W 74.5224, 27 May 2009,

L. Myers, B. Kondratieff, 2♂, 7♀ (CSUC); Essex Co.,

Seep to North Fork Boquet River flowing off Noble

Mountain, N 44.10820, W 73.69780, 9 May 2006, L.

Myers, 2♂, ♀ (CSUC); Pennsylvania, Center Co.,

Big Poe Creek, Poe Paddy Dr., N 40.82972, W

77.44472, 29 March 1995, E.C. Masteller, ♂, ♀; Erie

Co., Sixmile Creek, H and B Morgan, 7 April 1986,

E.C. Masteller, ♂ (CSUC); trib. to upper Fourmile

Creek, 27 April 1987, E.C. Masteller, 3♂, 9♀ (CSUC);

Fourmile Creek, Penn State Behrend Campus, N

42.12139, W 79.95416, 26 April 1981, E.C. Masteller,

♂, 3♀ (CSUC); Forest Co., Indian Spring, old

logging road, N 41.32791, 79.20204, 20 May 2009, S.

Harris, ♂ (CSUC).

Biological notes. The above records for S. amicalola

are the first of this genus from Georgia. The two

locations this species is known from are mid-

elevation streams (630–792 m) with nearby seeps.

Additional collections from these locations are

needed to confirm the identity of the female. Other

adult stoneflies collected with the holotype and

paratypes were Allocapnia aurora Ricker, 1952 and

Leuctra ferruginea (Walker, 1852).

New state records

Nemouridae

Amphinemura varshava (Ricker, 1952) was

described from near Warsaw in northern Indiana

and is a common inhabitant of cool streams in the

Midwest region having been reported from Indiana,

Illinois (DeWalt & Grubbs 2011), Kentucky (Ricker

1952), Ohio (DeWalt et al. 2012), and Wisconsin

(Grubbs et al. 2012). This species of

Amphinemurinae was collected near the edge of

Chattahoochee National Forest in the Ridge and

Valley Physiographic Province of Georgia. This

record constitutes a substantial southern range

extension.

Material examined. Georgia, Whitfield Co., Dalton

College Campus, [N 34.77335, W 85.00325], 29 April

1996, D. Yanega, ♂, ♀ (INHS).

Prostoia completa (Walker, 1852) was described

from Nova Scotia, Canada and has been reported

from every southeastern state except Florida and

Georgia (Grubbs et al. 2014). This species of

Nemourinae was collected from two locations in the

Piedmont Plateau Physiographic Province of

Georgia. Similar to the records plotted in James

(1972), these two localities represent the known

southern limit of the distribution of this widespread

species.

Material examined. Georgia, Putnam Co., Big

Indian Creek, Big Indian Creek Rd., N 33.39687, W

83.47433, 11 February 2017, C. Verdone, B.

Kondratieff, 5♂, 3♀ (CSUC); Murder Creek,

Hillsboro Rd., N 33.26861, W 83.49739, 11 February

2017, C. Verdone, B. Kondratieff, 5♂, 4♀, 3N

(CSUC).

Leuctridae

Leuctra carolinensis Claassen, 1923 was originally

described from Black Mountain, North Carolina and

has been reported from adjacent states of South

Carolina and Tennessee (Unzicker & McCaskill

1982, Stark et al. 1986, Kondratieff et al. 1995,

Stewart & Stark 2002). This species was collected

from the type locality of Remenus duffieldi, a 2nd order

stream in the Blue Ridge Physiographic Province of

Georgia.

Material examined. Georgia, Towns Co.,

Soapstone Creek, 2.9 km SE Brasstown Bald, N

Page 8: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 37

34.85588, W 83.78517, 17 May 2017, E.J. South, ♂

(INHS).

Leuctra ferruginea (Walker, 1852) is widespread in

the eastern United States and Atlantic Canada

(Harrison & Stark 2010). The only southeastern state

from which this species had yet to be reported was

Georgia. This species was collected from several

locations in the Blue Ridge and Piedmont Plateau

physiographic provinces of Georgia.

Material examined. Georgia, Dawson Co., Little

Amicalola Creek, Amicalola Falls State Park, N

34.57233, W 84.24107, 9 February 2017, C. Verdone,

B. Kondratieff, 3♂, 4♀ (CSUC); Fannin Co.,

[Hemptown Creek], 6 miles east of Morgantown,

Rt. 76, [N 34.88985, W 84.19029], 30 January 1963,

L.J. Stannard, W.S. Brooks, ♂ (INHS); Fulton Co.,

Fort McPherson, [N 33.7077, W 84.43565], 24 June

1943, H. Hoogstraal, ♂ (INHS); Gilmer Co., Tails

Creek, no location, 3 January 2013, C. Belcher ♂, ♀

(CSUC); Habersham Co., Tallulah River, Tallulah

Falls, [N 34.73056, W 83.39444], 10 May 1944, T. H.

Frison, H. H. Ross, ♂ (INHS); Lumpkin Co.,

Frogtown Creek, Hwy 19, Desoto Falls Rec. Area,

N 34.70649, W 83.91634, 10 February 2017, C.

Verdone, B. Kondratieff, 3♂, 4♀ (CSUC); Rabun

Co, Lake Burton, [N 34.79389, W 83.54056], 22 June

1940, H. H. Hobbs, ♂, 2♀ (INHS); Union Co., Wolf

Creek, Vogel State Park at Burnette Branch, N

34.76311, W 83.92679, 10 February 2017, C.

Verdone, B. Kondratieff, ♂, 2♀ (CSUC).

Leuctra sibleyi Claassen, 1923 is widely

distributed in the eastern United States and

Atlantic Canada, but has not been reported from

South Carolina, Georgia, or Florida. This species

occurs in small streams to medium-sized rivers

(Harper & Hynes 1971) and emerges in the spring

throughout its range. This species was collected

from two 3rd order streams in the Blue Ridge

Physiographic Province of northern Georgia.

Material examined. Georgia, Fannin Co.,

Noontootla Creek, 10.4 km NNE Amicalola Mtn. at

F.S. Rd. 58, N 34.68791, W 84.20182, 22 March 2016,

E.J. South, ♂ (INHS). Gilmer Co., Turniptown

Creek, US 76, [N 34.724009, W 84.43539], 23 April

2002, D. Etnier, ♂ (CSUC).

Leuctra tenella (Provancher, 1878) was originally

described from the vicinity of Quebec City, Quebec,

Canada. Ricker (1952) subsequently designated a

lectotype from Old Forge, New York. Leuctra tenella

is the sister species of L. carolinensis (Grubbs 2015)

and generally exhibits a more northeastern

distribution. However, this species has been

reported from Virginia (Kondratieff & Kirchner

1987), Great Smoky Mountain National Park in

North Carolina (Parker et al. 2007) and nearby

locations in Haywood and Macon counties (Grubbs

2015). This species was collected from a 3rd order

stream in the Blue Ridge Physiographic Province of

northern Georgia.

Material examined. Georgia, Union Co., Nottely

River, 8.0 km NNW Pinnacle Mountain, N 34.74962,

W 83.84646, 20 May 2017, E.J. South, ♂ (INHS).

Paraleuctra sara (Claassen, 1937) is the only species

of the genus that occurs in the eastern United States

(Stark & Kyzar 2001) and has been reported from

adjacent states of Alabama (Grubbs 2011), North

Carolina and South Carolina (Unzicker & McCaskill

1982, Stark et al. 1986, Stewart & Stark 1988,

Kondratieff et al. 1995). This species was collected

from a 3rd order stream in the Blue Ridge

Physiographic Province of northern Georgia.

Material examined. Fannin Co., Noontootla Creek,

10.4 km NNE Amicalola Mtn. at F.S. Rd. 58, N 34.68791,

W 84.20182, 22 March 2016, E.J. South, ♂, 2♀ (INHS).

Towns Co., trib. to Owl Creek, Rte. 180, Chattahoochee

National Forest, N 34.86261, W 83.76931, 7 March 2009,

S.A. Grubbs, ♂, ♀, (WKUC).

Zealeuctra fraxina Ricker & Ross, 1969 was

described from Kentucky and is one of three

Zealeuctra species that occurs in the Appalachian

Mountains and has been reported from adjacent

states of Tennessee and Alabama (Grubbs et al.

2013). This uncommon species was collected from

two streams in Chattahoochee National Forest in the

Ridge and Valley Physiographic Province of

northwestern Georgia.

Material examined. Georgia, Floyd Co., Johns

Creek, Everett Springs Rd., N 34.56416, W 85.10104,

8 February 2017, C. Verdone, B. Kondratieff, 4♂, ♀

(CSUC); Walker Co., trib. to Johns Creek, Keown

Falls Trail, N 34.6138, W 85.08757, 8 February 2017,

Page 9: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 38

C. Verdone, B. Kondratieff, ♂ (CSUC).

Capniidae

Allocapnia muskogee Grubbs & Sheldon, 2008 was

described from Cleburne and Clay counties

Alabama and included a paratype male specimen

from Lumpkin County, Georgia collected in 1964.

Through the courtesy of S.A. Grubbs an additional

male and three females of this uncommon Allocapnia

from Towns County are listed below. Both Georgia

locations are located in the Blue Ridge

Physiographic Province.

Material examined. Georgia, Towns Co., tributary

to High Shoals Creek, FR 283, Chattahoochee

National Forest, N 34.80912, W 83.72552, 7 March

2009, S.A. Grubbs, ♂, 3♀ (WKUC).

Allocapnia tsalagi Grubbs, 2008 was described

from a small spring in the Appalachian Plateau

Physiographic Province of northeastern Alabama.

This rare species was collected in the Ridge and

Valley Physiographic Province in northern Georgia

from a 1st order stream 122 km east, and nearly at

the same latitude as the type locality.

Material examined. Georgia, Murray Co., Head

Branch, Old Hwy 2, E of Cisco at Lawground

Branch, N 34.94523, W 84.72191, 2 January 2014, T.P.

Belcher, 7♂ (CSUC).

Allocapnia unzickeri Ross & Yamamoto, 1966 was

described from Tennessee and is believed to be

restricted to the Appalachian Plateau and

Cumberland River Valley (Ross & Ricker 1971). This

former Tennessee endemic was collected from a 3rd

order stream in the Appalachian Plateau

Physiographic Province in the northwest corner of

Georgia, 16.9 km south of the Tennessee border.

Material examined. Georgia, Dade Co., Bear Creek,

SR 189, Cloudland State Park, N 34.82823, W

85.45934, 8 February 2017, C. Verdone,B.

Kondratieff, ♂, 2♀ (CSUC).

Nemocapnia carolina Banks, 1938 has been

reported from Quebec, Canada to Florida and west

to central Arkansas (Stark et al. 2016). This species

has been reported from the Wabash River drainage

of Illinois and Indiana (Frison 1935), but is thought

to be extirpated from those states (DeWalt & Grubbs

2011). Nemocapnia carolina has been reported from

several counties in adjacent states near the Georgia

state line, but has not been formally reported from

Georgia (Stark et al. 2016). This uncommon winter

stonefly was collected from small and large order

streams in the Piedmont Plateau and Coastal Plain

physiographic provinces of Georgia.

Material examined. Georgia, Bleckley Co.,

Ocmulgee River, James Dykes Park, N 32.41439, W

83.48319, 12 February 2017 C. Verdone, B.

Kondratieff, 20♂, 14♀ (CSUC); Greene Co., Harris

Creek, Nichols Rd., N 33.72509, W 83.23737, 11

February 2017, C. Verdone, B. Kondratieff, ♂, ♀

(CSUC); Oconee River, Hwy 15, Oconee National

Forest campground, N 33.72149, W 83.29095, 11

February 2017, C. Verdone, B. Kondratieff, 20♂, 8♀

(CSUC); Houston Co., Burnham Brand Creek, Hwy

26, SE of Elko, N 32.31585, W 83.66442, 12 February

2017, C. Verdone, B. Kondratieff, ♀ (CSUC); Macon

Co., Flint River, Montezuma Bluffs Park, N

32.33682, W 84.03118, 12 February 2017, C. Verdone,

B. Kondratieff, 5♂, 5♀ (CSUC); Putnam Co., Little

River, Glades Rd., N 33.37238 W 83.4773, 11

February 2017, C. Verdone, B. Kondratieff, 3♂, 2♀

(CSUC); Stewart Co., Hannahatchee Creek, Co. Rd.

150, N 32.14157, W 84.75333, 12 February 2017, C.

Verdone, B. Kondratieff, 3♂, 3♀ (CSUC).

Paracapnia angulata (Hanson, 1961) is not

common in the southern Appalachians, but is the

most widespread Nearctic species in the genus. It

has been recorded from Atlantic Canada to

Arkansas and as far west as Colorado (Stark &

Baumann 2004). This species has been reported from

Macon, Jackson and Transylvania counties in North

Carolina near the Georgia state line (Beaty 2015).

The records presented here were collected over 50

years ago, but were never reported.

Material examined. Georgia, Fannin Co.,

[Hemptown Creek], 6 miles east of Morgantown, Rt.

76, [N 34.88985, W 84.19029], 30 January 1963, L.J.

Stannard, W.S. Brooks, 4♂ (INHS); Union Co.,

[Youngcane Creek], 7 mi west of Blairsville, Rt. 76,

[N 34.83568, W 84.08412], 30 January 1963, L.J.

Stannard, W.S. Brooks, 2♂, ♀ (INHS).

Taeniopterygidae

Page 10: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 39

Oemopteryx contorta (Needham & Claassen, 1925)

was described from Jaffery, New Hampshire and

has been reported from adjacent states of Alabama

(Grubbs 2006), Tennessee (Nelson 1982), and North

Carolina (Kondratieff et al. 1995). This species of

Brachypterainae was collected from a 2nd order

stream in the Appalachian Plateau Physiographic

Province in the northwest corner of Georgia.

Material examined. Georgia, Dade Co., Daniel

Creek, Cloudland State Park, N 34.82457, W

85.49074, 8 February 2017, C. Verdone, B.

Kondratieff, 2♂, 4♀, 2N (CSUC).

Strophopteryx appalachia Ross & Ricker, 1975 was

originally described from Virginia and has been

reported from adjacent states of North Carolina

(Kondratieff et al. 1995), Tennessee, and South

Carolina (Unzicker & McCaskill 1982 Stark et al.

1986, Stewart & Stark 1988, Kondratieff et al. 1995,).

This species of Brachypterainae was collected from

a 2nd order stream in the Piedmont Plateau

Physiographic Province, 51.5 km northwest of

Atlanta, Georgia and the Chattooga River at the

South Carolina border.

Material examined. Georgia, Paulding Co., Bluffy

Creek, Hulsey Town Rd., N 33.8926, W 84.92486, 8

February 2017, C. Verdone, B. Kondratieff, ♂, 1N

(CSUC). Rabun Co., Chattooga River, Hwy 76

Fishing Access, N 34.81404, W 83.30647, 10 February

2017, C. Verdone, B. Kondratieff, ♀ (CSUC).

Strophopteryx fasciata (Burmeister, 1839) is

widespread in the eastern United States and

Atlantic Canada (DeWalt et al. 2017). This species

has been reported from adjacent states of Alabama

(Grubbs 2011), North Carolina and South Carolina

(Unzicker & McCaskill 1982, Stark et al. 1986,

Stewart & Stark 1988, Kondratieff et al. 1995). This

species of Brachypterainae was collected from

several locations in the Blue Ridge and Piedmont

Plateau physiographic provinces of Georgia.

Material examined. Georgia, Murray Co.,

Conasauga River, Old Hwy 2, N 34.97461, W

84.64487, 9 February 2017, C. Verdone, B.

Kondratieff, 2♂, 3♀ (CSUC); Holly Creek, Camp Rd.

Day Use Area, N 34.81181, W 84.6608, 9 February

2017, C. Verdone, B. Kondratieff, ♂ (CSUC); Pickens

Co., Talking Rock Creek, 6.3 km WNW Talking

Rock at GA-136, N 34.52624, W 84.57093, 18

February 2017, E.J. South, ♂ (INHS); Putnam Co.,

Big Indian Creek, Big Indian Creek Rd., N 33.39687,

W 83.47433, 11 February 2017, C. Verdone, B.

Kondratieff, 2♂ (CSUC); Murder Creek, Hillsboro

Rd., N 33.26861, W 83.49739, 11 February 2017, C.

Verdone, B. Kondratieff, 4♂, 3♀, 5N (CSUC).

Chloroperlidae

Sweltsa mediana (Banks, 1911) was described from

Black Mountain, North Carolina and has been

subsequently reported from southwestern Virginia

(Surdick 2004) to northwestern South Carolina

(Grubbs 2011). Adults are typically active from late

April to early July (Surdick 2004). This species of

Chloroperlinae was collected from a 2nd order

stream in the Ridge and Valley Physiographic

Province of northern Georgia.

Material examined. Georgia, Murray Co., Rock

Creek, US 411, [N 34.69271, W 84.73510], 24 April

1999, D.A. Etnier, B. Haue, ♂ (CSUC).

Perlidae

Agnetina annulipes (Hagen, 1861) has been

reported from Ohio (DeWalt et al. 2012), southern

Indiana (DeWalt & Grubbs 2011), and from

Pennsylvania to Florida west to Louisiana (Stark

1986). Stark (1986) indicated that the distribution of

this species was similar to Acroneuria arenosa (Pictet,

1841) and Paragnetina fumosa (Banks, 1902), both of

which have been reported from Georgia (Ricker

1949, Stark & Gaufin 1976). Specimens presented

here were collected nearly 80 years ago by the late

Dr. P.W. Fattig. Described as a “tireless collector”

(Lund 1954) the former Emory University

entomologist made many contributions to the

documentation of various insect groups from

Georgia.

Material examined. Georgia, Upson Co., [Flint

River], Whispering Pines Tourism Camp, 13 mi. S

Thomaston, [N 32.72215, W 84.23258], 8 June 1939,

P.W. Fattig, 4♂, 3♀ (INHS).

Neoperla coosa Smith & Stark, 1998 was described

from the Coosa River drainage in Chilton County,

Page 11: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 40

Alabama. This species has since been reported from

adjacent states of North Carolina (Lenat et al. 2009)

and Tennessee (DeWalt & Heinold 2005). This

species of Perlinae was collected in the Piedmont

Plateau Physiographic Province of Georgia near the

Fall Line, the escarpment separating the Piedmont

Plateau from the Coastal Plain.

Material examined. Georgia, Upson Co., [Flint

River], Whispering Pines Tourism Camp 13 mi. S

Thomaston, [N 32.72215, W 84.23258], 8 June 1939,

P.W. Fattig, 5♂, 3♀ (INHS).

Neoperla stewarti Stark & Baumann, 1978 is

widely distributed in the southeastern and

midwestern United States and has been reported

from adjacent states of Alabama, Tennessee (Stark

1990) and North Carolina (Lenat et al. 2009). This

species was collected from a low elevation 4th order

stream in the Ridge and Valley Physiographic

Province of northwestern Georgia.

Material examined. Georgia, Catoosa Co.,

Chickamauga Creek, Ringgold, [N 34.91525, W

85.12374], 14 June 1939, P.W. Fattig, ♂, ♀ (INHS).

Perlesta decipiens (Walsh, 1862) is the most

widespread Nearctic species in the genus, occurring

from the sky islands of New Mexico and Arizona

(Cary & Jacobi 2008) to east of the Appalachian

Mountains in Virginia (Stark 1989). This species was

collected from a low elevation 4th order stream in the

Ridge and Valley Physiographic Province of

northwestern Georgia.

Material examined. Georgia, Catoosa Co.,

Chickamauga Creek, Ringgold, [N 34.91525, W

85.12374], 14 June 1939, P.W. Fattig, 5♂, ♀ (INHS).

Perlesta ephelida Grubbs & DeWalt, 2012 is widely

distributed across the central and eastern United

States and frequently occurs in lower elevation mid-

sized streams (Grubbs & DeWalt 2012). This species

was collected from a low elevation 4th order stream

in the Ridge and Valley Physiographic Province of

northwestern Georgia and an unspecified location

near Ball Ground in the Piedmont Plateau

Physiographic Province.

Material examined. Georgia, Catoosa Co.,

Chickamauga Creek, Ringgold, [N 34.91525, W

85.12374], 14 June 1939, P.W. Fattig, 6♂, 7♀ (INHS);

Cherokee Co., Ball Ground [N 34.33727, W

84.37795], 27 June 1932, P.W. Fattig, ♂ (INHS).

Perlesta lagoi Stark, 1989 was originally described

from Hinds County, Mississippi. This species has

since been reported from adjacent states of Alabama

(Grubbs 2012) and Tennessee (Kondratieff &

Kirchner 2002). This species was collected from a

low elevation 3rd order stream in the upper Coastal

Plain Physiographic Province of Georgia.

Material examined. Georgia, Crawford Co.,

Culpepper Creek, 5 mi. SE Roberta, [N 32.69165, W

84.00792], 5 May 1939, P.W. Fattig, 2♂, 3♀ (INHS).

Perlesta ouabache Grubbs & DeWalt, 2011 is

known only from Indiana. A single male specimen

from an unknown location near Ball Ground,

Georgia was examined that possesses characters

consistent with P. ouabache. However, because of the

disjunction of this record from the known

distribution, fresh material is needed to confirm the

presence of this species in Georgia.

Material examined. Georgia, Cherokee Co., Ball

Ground [N 34.33727, W 84.37795], 27 June 1932,

P.W. Fattig, ♂ (INHS).

Perlesta shawnee Grubbs & Stark, 2004 was

described from Illinois and Indiana and has

subsequently been reported from Alabama, North

Carolina, Virginia (Grubbs & DeWalt 2008) and

Kentucky (Grubbs 2012). This species was collected

from two locations in the Piedmont Plateau

Physiographic Province of Georgia.

Material examined. Georgia, Cherokee Co., Ball

Ground [N 34.33727, W 84.37795], 27 June 1932,

P.W. Fattig, 4♂ (INHS); Fulton Co., Fort McPherson,

[N 33.7077, W 84.43565], 3 June 1943, H. Hoogstraal,

♂ (INHS); same location, 30 July 1943, H.

Hoogstraal, 3♂ (INHS).

Perlodidae

Cultus verticalis (Banks, 1920) has been reported

from Atlantic Canada to Tennessee and North

Carolina (Stark et al. 1988). This species often occurs

in small to medium sized streams in the

Appalachians (Myers & Kondratieff 2009). It is

Page 12: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 41

reported from a 3rd order stream in the Blue Ridge

Physiographic Province of northern Georgia.

Material examined. Georgia, Fannin Co.,

Noontootla Creek, 10.4 km NNE Amicalola Mtn. at

F.S. Rd. 58, N 34.68791, W 84.20182, 8 May 2016, E.J.

South, ♂ (INHS).

Table 2 presents 100 species for Georgia and should

be considered a work in progress as there are several

genera that are likely underrepresented in our

records, namely Leuctra Stephens, Isoperla Banks,

Neoperla Needham and Perlesta Banks.

DISCUSSION

During this study, a male specimen of Perlesta,

close to P. xube Stark & Rhodes, 1997 and several

males and females of Agnetina Klapálek, could not

be satisfactorily determined to species. Additional

properly prepared adult specimens of these

stoneflies are required for specific determination.

Perlesta in particular, must be collected alive and the

aedeagus fully everted to allow for specific

determination. Briefly, we (CJV & BCK) collected

live specimens in the field, brought them back to our

lodging, and prepared these specimens under a

dissecting microscope. Using wide-tipped forceps,

we gently squeezed the aedeagus out to near its full

extent, then submerged the specimen in near boiling

water to fix the aedeagus in place. Often, some

additional massaging of these specimens was

required to more fully evert the terminal filament or

dorsal caecum. Specimens were then preserved in

80% ethanol. Our coauthors employ slightly

different methods, but the one described above

worked well.

Stonefly species richness in the eastern United

States has been attributed to the heterogeneity of

lotic habitats associated with topography and the

five major physiographic provinces (Kondratieff &

Kirchner 1987, Kondratieff et al. 2017): Appalachian

Plateau, Blue Ridge, Coastal Plain, Piedmont

Plateau, and Ridge and Valley (Woodward &

Hoffman 1991). Many of the conditions that have

resulted in high species diversity in other eastern

states are present in Georgia including all five

physiographic provinces and relatively high

topographic relief, which ranges from sea level to

1458 m (4784 ft.) at Brasstown Bald. Georgia is a

state with potential for several additional new state

records and undescribed species. The mountainous

regions have been relatively well collected, but the

Piedmont and Coastal Plain physiographic

provinces, which cover the majority of the state,

remain poorly surveyed.

Several of the species presented as new state

records and in the checklist are historical. Local and

statewide extirpation of plecopteran species have

been documented in Illinois (DeWalt et al. 2005,

DeWalt & Grubbs 2011), Indiana (DeWalt & Grubbs

2011), Michigan (Grubbs et al. 2012) and Ohio

(DeWalt et al. 2012) As such, modern collections are

needed to confirm the continued presence of species

in this imperiled insect order. Entomologists

interested in collecting stoneflies in Georgia are

required to obtain collecting/research permits from

the following agencies: United States Forest Service,

Georgia Department of Natural Resources Parks,

Recreation and Historic Sites Division, and Georgia

Department of Natural Resources Law Enforcement

Division.

Page 13: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 42

Table 2. List of Georgia stoneflies.

Family Species First Reference

Nemouridae Amphinemura appalachia Baumann, 1996 Baumann (1996)

Amphinemura delosa (Ricker, 1952) Ricker (1952)

Amphinemura nigritta (Provancher, 1876) Ricker (1952) Amphinemura varshava (Ricker, 1952) new state record

Amphinemura wui (Claassen, 1936) Ricker (1952)

Prostoia completa (Walker, 1852) new state record

Prostoia hallasi Kondratieff & Kirchner, 1984 Grubbs et al. (2014)

Soyedina amicalola Verdone & Kondratieff sp. n. Leuctridae Leuctra alexanderi Hanson, 1941 Kondratieff & Nelson (1995)

Leuctra biloba Claassen, 1923 Claassen (1923) Leuctra carolinensis Claassen, 1923 new state record

Leuctra ferruginea (Walker, 1852) new state record

Leuctra moha Ricker, 1952 Ricker (1952) Leuctra sibleyi Claassen, 1923 new state record Leuctra tenella (Provancher, 1878) new state record Paraleuctra sara (Claassen, 1937) new state record Zealeuctra fraxina Ricker & Ross, 1969 new state record

Capniidae Allocapnia aurora Ricker, 1952 Ricker (1952) Allocapnia muskogee Grubbs & Sheldon, 2008 Grubbs & Sheldon 2008

Allocapnia mystica Frison, 1929 Ross & Ricker (1971)

Allocapnia recta (Claassen, 1924) Ross & Ricker (1971)

Allocapnia rickeri Frison, 1942 Ross & Ricker (1971)

Allocapnia tsalagi Grubbs, 2008 new state record

Allocapnia unzickeri Ross & Yamamoto, 1966 new state record

Allocapnia virginiana Frison, 1942 Ross & Ricker (1971)

Allocapnia wrayi Ross, 1964 Ross & Ricker (1971)

Nemocapnia carolina Banks, 1938 new state record Paracapnia angulata (Hanson, 1961) new state record Taeniopterygidae Oemopteryx contorta (Needham & Claassen, 1925) new state record

Strophopteryx appalachia Ross & Ricker, 1975 new state record

Strophopteryx fasciata (Burmeister 1839) new state record

Taeniopteryx burksi Ricker & Ross, 1968 Ricker & Ross (1968)

Taeniopteryx lonicera Ricker & Ross, 1968 Ricker & Ross (1968)

Taeniopteryx maura (Pictet, 1841) Ricker & Ross (1968)

Taeniopteryx parvula Banks, 1918 Ricker & Ross (1968)

Taeniopteryx ugola Ricker & Ross, 1968 Fullington & Stewart (1980) Chloroperlidae Alloperla atlantica Baumann, 1974 Baumann (1974)

Alloperla chloris Frison, 1934 Surdick (1985)

Alloperla idei (Ricker, 1935) Surdick (1985)

Alloperla nanina Banks, 1911 Needham & Claassen (1925)

Page 14: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 43

Alloperla petasata Surdick, 2004 Surdick (2004)

Alloperla usa Ricker, 1952 Ricker (1952)

Haploperla brevis (Banks, 1895) Needham & Claassen (1925)

Suwallia marginata (Banks, 1897) Surdick (1985)

Sweltsa lateralis (Banks, 1911) Needham & Claassen (1925) Sweltsa mediana (Banks, 1911) new state record

Sweltsa voshelli Kondratieff & Kirchner, 1991 Surdick (2004) Perlidae Acroneuria abnormis (Newman, 1838) Berner (1948)

Acroneuria arenosa (Pictet, 1841) Stark & Gaufin (1976)

Acroneuria arida (Hagen, 1861) Frison (1942)

Acroneuria carolinensis (Banks, 1905) Stark (2004)

Acroneuria evoluta Klapálek, 1909 Frison (1942)

Acroneuria filicis Frison, 1942 Stark & Gaufin (1976)

Acroneuria perplexa Frison, 1937 Stark & Gaufin (1976)

Acroneuria petersi Stark & Gaufin, 1976 Stark & Gaufin (1976) Agnetina annulipes (Hagen, 1861) new state record

Agnetina flavescens (Walsh, 1862) Stark (1986)

Attaneuria ruralis (Hagen, 1861) Berner (1948)

Beloneuria georgiana (Banks, 1914) Banks (1914)

Beloneuria stewarti Stark & Szczytko, 1976 Stewart & Stark (1993)

Eccoptura xanthenes (Newman, 1838) Newman (1838)

Neoperla clymene (Newman, 1839) Newman (1839) Neoperla coosa Smith & Stark, 1998 new state record Neoperla stewarti Stark & Baumann, 1978 new state record

Paragnetina fumosa (Banks, 1902) Ricker (1949)

Paragnetina immarginata (Say, 1823) Needham & Claassen (1925)

Paragnetina kansensis (Banks, 1905) Berner (1948) Perlesta decipiens (Walsh, 1862) new state record Perlesta ephelida Grubbs & DeWalt, 2012 new state record Perlesta lagoi Stark, 1989 new state record Perlesta placida (Hagen, 1861) Berner (1948)

Perlesta shawnee Grubbs & Stark, 2004 new state record Perlinella drymo (Newman, 1839) Newman (1839)

Perlinella ephyre (Newman, 1839) Newman (1839) Perlodidae Clioperla clio (Newman, 1839) Newman (1839)

Cultus decisus isolatus (Banks, 1920) Ricker (1952)

Cultus verticalis (Banks, 1920) new state record Diploperla duplicata (Banks, 1920) Ricker (1952)

Helopicus bogaloosa Stark & Ray, 1983 Stark & Ray (1983)

Helopicus subvarians (Banks, 1920) Ricker (1952)

Isoperla bellona Banks, 1911 Needham & Claassen (1925)

Isoperla chickamauga Szczytko & Kondratieff, 2015 Szczytko & Kondratieff (2015)

Isoperla dicala Frison, 1942 Szczytko & Kondratieff (2015)

Page 15: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 44

Isoperla frisoni Illies, 1966 Szczytko & Kondratieff (2015)

Isoperla holochlora Klapálek, 1923 Ricker (1938)

Isoperla lenati Szczytko & Kondratieff, 2015 Szczytko & Kondratieff (2015)

Isoperla nelsoni Szczytko & Kondratieff, 2015 Szczytko & Kondratieff (2015)

Malirekus hastatus (Banks, 1920) Ricker (1952)

Remenus bilobatus (Needham & Claassen, 1925) Ricker (1952)

Remenus duffieldi Nelson & Kondratieff, 1995 Kondratieff & Nelson (1995)

Yugus arinus (Frison, 1942) Ricker (1952)

Yugus bulbosus (Frison, 1942) Ricker (1952) Peltoperlidae Tallaperla anna (Needham & Smith, 1916) Needham & Smith (1916)

Tallaperla cornelia (Needham & Smith, 1916) Needham & Smith (1916)

Tallaperla laurie (Ricker, 1952) Ricker (1952)

Tallaperla maria (Needham & Smith, 1916) Ricker (1952)

Viehoperla ada (Needham & Smith, 1916) Needham & Smith (1916) Pteronarcyidae Pteronarcys biloba Newman, 1838 Ricker (1952)

Pteronarcys dorsata (Say, 1823) Nelson & Hanson (1971)

Pteronarcys scotti Ricker, 1952 Ricker (1952)

ACKNOWLEDGEMENTS

We thank Chip Belcher for providing the

Allocapnia tsalagi record, Dr. Mike Ferro, Clemson

University, Clemson, South Carolina and Dr. Oliver

Flint, Smithsonian Institution, Washington, DC, for

loaning specimens, the United States Forest Service

and the Georgia Department of Natural Resources

for providing the necessary permits required to

collect insects in the state of Georgia. Dr. Scott

Grubbs and an anonymous reviewer are thanked

for providing helpful comments that improved the

manuscript. We especially thank Mike Kippenhan,

Portland, Oregon, for preparing the illustrations.

REFERENCES

Banks N. 1914. New neuropteroid insects, native

and exotic. Proceedings of the Academy of

Natural Sciences of Philadelphia, 1:608–632.

Baumann, R.W. 1974. What is Alloperla imbecilla

(Say)? Designation of a neotype, and a new

Alloperla from eastern North America

(Plecoptera: Chloroperlidae). Proceedings of the

Entomological Society of Washington, 87:257–

264.

Baumann, R.W. & S.A. Grubbs. 1996. Two new

species of Soyedina (Plecoptera: Nemouridae)

from the Appalachian Mountains.

Entomological News, 107(4):220–224.

http://www.biodiversitylibrary.org/page/2700498

Beaty, S.R. 2015. The Plecoptera of North Carolina:

A biologist’s handbook for the identification of

stonefly nymphs with standard taxonomic effort

levels. Version 4.0. North Carolina Department

of Environmental Quality, Division of Water

Resources, Biological Assessment Branch.

Raleigh, North Carolina. iv + 91 pp.

https://ncdenr.s3.amazonaws.com/s3fs-public/Water

Quality/Environmental Sciences/BAU/Benthos

Reference/BAB Taxonomy Doc - Plecoptera –

version 204.1- complete.pdf

Berner, L. 1948. Records of stoneflies from Florida

(Plecoptera). The Florida Entomologist, 31(1):21–

23.

Cary, S.J. & G.Z. Jacobi. 2008. Zoogeographic

affinities of southwestern USA Plecoptera.

Pp.133–157. In: International advances in the

ecology, zoogeography, and systematics of

mayflies and stoneflies. F.R. Hauer, J.A. Stanford

& R.L. Newell (Eds.) University of California

Press, 422 pp.

Claassen, P.W. 1923. New species of North

American Plecoptera. The Canadian

Entomologist, 55(12):281–292.

DeWalt, R.E. & B.D. Heinold. 2005. Summer

emerging Ephemeroptera, Plecoptera, and

Trichoptera of Abrams Creek, Great Smoky

Page 16: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 45

Mountains National Park. Proceedings of the

Entomological Society of Washington, 107(1):34–

48. http://www.biodiversitylibrary.org/page/32142754

DeWalt, R.E. & S.A. Grubbs. 2011. Updates to the

stonefly fauna of Illinois and Indiana. Illiesia,

7(3):31–50. http://illiesia.speciesfile.org/papers/

Illiesia07-03.pdf

DeWalt, R.E., C. Favret & D.W. Webb. 2005. Just

how imperiled are aquatic insects? A case study

of stoneflies (Plecoptera) in Illinois. Annals of the

Entomological Society of America, 98(6):941–950.

DeWalt, R.E., M.D. Maehr, U. Neu-Becker & G.

Stueber. 2017. Plecoptera Species File Online.

Version5.0/5.0. http://Plecoptera.SpeciesFile.org/

(Accessed 24 February 2017)

DeWalt, R.E., Y. Cao, T. Tweddale, S. Grubbs, L.

Hinz, M. Pessino & J. Robinson. 2012. Ohio USA

stoneflies (Insecta, Plecoptera): species richness

estimation, distribution of functional niche traits,

drainage affiliations, and relationships to other

states. ZooKeys, 178, p.1.

https://www.ncbi.nlm.nih.gov/pmc/articles/PM

C3317619/pdf/ZooKeys-178-001.pdf

Frison, T.H. 1935. The stoneflies, or Plecoptera of

Illinois. Illinois Natural History Survey Bulletin,

20:281–467. http://www.nativefishlab.net/library

/textpdf/18843.pdf

Frison, T.H. 1942. Studies of North American

Plecoptera with special reference to the fauna of

Illinois. Illinois Natural History Survey Bulletin,

21:78–99.

Fullington, K.E. & K.W. Stewart. 1980. Nymphs of

the stonefly genus Taeniopteryx (Plecoptera:

Taeniopterygidae) of North America. Journal of

the Kansas Entomological Society, 53:237–259.

Grubbs, S.A. 2006. Allocapnia sano, a new species of

snowfly (Plecoptera: Capniidae) from Alabama,

USA, plus six new state records. Zootaxa,

1197(1):39–43. https://www.researchgate.net/

profile/Scott_Grubbs/publication/228919016

Grubbs, S.A. 2011. A review of stonefly (Insecta;

Plecoptera) taxonomic research in Alabama, with

new state records and an updated checklist.

Illiesia, 7(2):24–30. http://illiesia.speciesfile.org/

papers/Illiesia07-02.pdf

Grubbs, S.A. 2012. An amended redescription of the

male of Perlesta etnieri (Plecoptera: Perlidae) plus

new Perlesta state records. Illiesia, 8(9):111–113.

http://illiesia.speciesfile.org/papers/Illiesia08-09.pdf

Grubbs, S.A. 2015. Leuctra schusteri, a new stonefly

species (Plecoptera: Leuctridae) of the L. tenuis

(Pictet) group from the southeastern USA.

Illiesia, 11(12):147–166.

http://illiesia.speciesfile.org/papers/Illiesia11-12.pdf

Grubbs, S.A. & A.L. Sheldon. 2008. Allocapnia

muskogee and A. menawa, new species of

snowflies (Plecoptera: Capniidae) from the

Talladega National Forest region of eastern

Alabama, USA, plus four new state records.

Illiesia, 4(11):99–109. http://illiesia.speciesfile.org

/papers/Illiesia04-11.pdf

Grubbs, S.A. & R.E. DeWalt. 2008. Taxonomic and

distributional notes on Perlesta teaysia, P.

golconda, and P. shawnee (Plecoptera: Perlidae).

Illiesia, 4(14):143–149. Illiesia, 4(14):143–149.

http://illiesia.speciesfile.org/papers/Illiesia04-14.pdf

Grubbs S.A. & R.E. DeWalt. 2012. Perlesta ephelida, a

new Nearctic stonefly species (Plecoptera,

Perlidae). ZooKeys 194: 1–15. doi:

10.3897/zookeys.194.2972

Grubbs, S.A., B.C. Kondratieff, B.P. Stark & R.E.

DeWalt. 2013. A review of the Nearctic genus

Zealeuctra Ricker (Plecoptera, Leuctridae), with

the description of a new species from the

Cumberland Plateau region of eastern North

America. ZooKeys, 344:17–47.

https://doi.org/10.3897/zookeys.344.5912

Grubbs, S.A., M. Pessino & R.E. DeWalt. 2012.

Michigan Plecoptera (stoneflies): distribution

patterns and an updated species list. Illiesia,

8(18):162–173. http://illiesia.speciesfile.org/papers/

Illiesia08-18.pdf

Grubbs, S.A., R.W. Baumann, R.E. DeWalt & T.

Tweddale. 2014. A review of the Nearctic genus

Prostoia (Ricker) (Plecoptera, Nemouridae), with

the description of a new species and a surprising

range extension for P. hallasi Kondratieff &

Kirchner. ZooKeys, 401:11–30.

https://doi.org/10.3897/zookeys.401.7299

Harper, P.P. & H.B.N. Hynes. 1971. The Leuctridae

of eastern Canada (Insecta; Plecoptera).

Canadian Journal of Zoology, 49(6):915–920.

http://www.nrcresearchpress.com/doi/pdf/10.11

39/z71-136

Harrison, A.B. & B.P. Stark. 2010. Two new species

of stoneflies in the Leuctra ferruginea group

Page 17: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 46

(Plecoptera: Leuctridae), with notes on the

Leuctra species known for Mississippi and

Alabama, U.S.A. Illiesia, 6(03):16-33.

http://illiesia.speciesfile.org/papers/Illiesia06-03.pdf

James, A.M. 1972. The stoneflies (Plecoptera) of

Alabama. Unpublished Doctoral dissertation,

Auburn University, Auburn, Alabama. 169 pp.

Kondratieff, B.C. & C.H. Nelson. 1995. A review of

the genus Remenus Ricker (Plecoptera:

Perlodidae), with the description of two new

species. Proceedings of the Entomological

Society of Washington, 97(3): 596–602.

Kondratieff, B.C. & R.F. Kirchner. 1987. Additions,

taxonomic corrections, and faunal affinities of

the stoneflies (Plecoptera) of Virginia, USA.

Proceedings of the Entomological Society of

Washington, 89:24–30.

http://www.biodiversitylibrary.org/page/16262424

Kondratieff, B.C. & R.F. Kirchner. 2002. Perlesta

etnieri (Plecoptera: Perlidae), a new species of

stonefly from Tennessee. Proceedings of the

Entomological Society of Washington, 104:51–55.

http://www.biodiversitylibrary.org/page/16173598

Kondratieff, B.C., C.J. Verdone & S. Roble. 2017.

New records of stoneflies (Plecoptera) from

Virginia, U.S.A. Perla, 35:22–27.

http://plecoptera.speciesfile.org/HomePage/Plec

optera/LitArchive/PerlaNo35_2017.pdf

Kondratieff, B.C., R.F. Kirchner & D.R. Lenat. 1995.

A review of stonefly records (Plecoptera:

Hexapoda) of North Carolina and South

Carolina. Brimleyana, (23):25–40.

https://www.researchgate.net/publication/26763

1352_A_Review_of_Stonefly_Records_Plecopter

a_Hexapoda_of_North_Carolina_and_South_C

arolina

Lenat, D.R., R.E. Zuellig B.C. Kondratieff & S.R.

Beaty. 2009. Distribution of Neoperla (Plecoptera:

Perlidae) in North Carolina, with new state

records for three species. Illiesia, 5(15):164–168.

http://illiesia.speciesfile.org/papers/Illiesia05-

15.pdf

Lund, H.O. 1954. Perry Wilbur Fattig 1881–1953.

Journal of Economic Entomology, 47(2):352.

https://doi.org/10.1093/jee/47.2.352

Myers, L.W. & B.C. Kondratieff. 2009. Descriptions

of the nymphs of eastern North Americans of

Cultus (Plecoptera: Perlodidae). New York

Entomological Society, 115(2):109–114.

Needham, J.G. & L.W. Smith. 1916. The stoneflies of

the genus Peltoperla. The Canadian Entomologist,

48(3):80–88.

Needham, J.G. & P.W. Claassen. 1925. Monograph

of the Plecoptera or stoneflies of America, north

of Mexico. Entomological Society of America,

Thomas Say Foundation, 2:1–397.

Nelson, C.H. 1982. Notes on the life histories of

Strophopteryx limata (Frison) and Oemopteryx

contorta (Needham and Claassen) (Plecoptera:

Taeniopterygidae) in Tennessee. Journal of the

Tennessee Academy of Science, 57:9–15.

Nelson, C.H. & J.F. Hanson. 1971. Contribution to

the anatomy and phylogeny of the family

Pteronarcyidae (Plecoptera). Transactions of the

American Entomological Society, 97(1):123–200.

Newman, E. 1838. Entomological notes. The

Magazine of Natural History, 5:168–181, 372–

402, 483–500.

Newman, E. 1839. On the synonymy of the Perlites,

together with brief characters of the old, and of a

few new species. The Magazine of Natural

History, 3:32–37, 84–90.

Parker, C.R., O.S. Flint Jr., L.M. Jacobus, B.C.

Kondratieff, W.P. McCafferty & J.C. Morse. 2007.

Ephemeroptera, Plecoptera, Megaloptera, and

Trichoptera of Great Smoky Mountains National

Park. Southeastern Naturalist, 6(sp2):159–174.

https://www.jstor.org/stable/pdf/4541006.pdf

Ricker, W.E. 1938. Notes on specimens of American

Plecoptera in European collections. Transactions

of the Royal Canadian Institute, 22(1):129-156.

Ricker, W.E. 1949. The North American species of

Paragnetina (Plecoptera, Perlidae). Annals of the

Entomological Society of America, 42(3):279–288.

https://doi.org/10.1093/aesa/42.3.279

Ricker, W.E. 1952. Systematic studies of Plecoptera.

Indiana University Publications Science Series,

18:1–200. http://www.nativefishlab.net/library/

textpdf/16861.pdf

Ricker, W.E. & H.H. Ross. 1968. North American

species of Taeniopteryx (Plecoptera, Insecta).

Journal of the Fisheries Board of Canada, 25(7):

1423–1439.

Ross, H.H. & W.E. Ricker. 1971. The classification,

evolution, and dispersal of the winter stonefly

genus Allocapnia. Illinois Biological Monographs

Page 18: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 47

45. University of Illinois Press, Urbana, Illinois.

166 pp. http://www.ideals.illinois.edu/bitstream

/handle/2142/27332/classificationev45ross.pdf

Stark, B.P. 1986. The Nearctic species of Agnetina

(Plecoptera: Perlidae). Journal of the Kansas

Entomological Society, 59:437–445.

http://www.jstor.org/stable/pdf/25084804.pdf

Stark, B.P. 1989. Perlesta placida (Hagen), an eastern

Nearctic species complex (Plecoptera: Perlidae).

Insect Systematics & Evolution, 20(3):263–286.

Stark, B.P. 1990. Neoperla clymene revisited:

systematics of the Nearctic species complexes

(Plecoptera: Perlidae). Pp. 299–310. In: Mayflies

and stoneflies life histories and biology:

proceedings of the 5th International

Ephemeroptera Conference and the 9th

International Plecoptera Conference. Campbell,

I.C. (Ed.) (Vol. 44). Kluwer Academic Publishers,

366 pp. https://link.springer.com/content/pdf/10.1007%

2F978-94-009-2397-3.pdf

Stark, B.P. 2004. Perlidae (The Stones) Pp. 61–148. In:

Stoneflies (Plecoptera) of eastern North America,

Volume II, Chloroperlidae, Perlidae, and

Perlodidae (Perlodinae). B.P. Stark, B.J. Armitage

(Eds.) Bulletin of the Ohio Biological Survey,

Columbus, Ohio. 192 pp.

Stark, B.P., A.B. Harrison & K.C. Nye. 2016. Records

of an uncommon stonefly, Nemocapnia carolina

Banks (Plecoptera: Capniidae), in South Carolina

and nearby states. Illiesia, 12(02):10–14.

http://illiesia.speciesfile.org/papers/Illiesia12-02.pdf

Stark, B.P. & A.R. Gaufin. 1976. The Nearctic species

of Acroneuria (Plecoptera: Perlidae). Journal of

the Kansas Entomological Society, 49(2):221–253.

http://www.jstor.org/stable/pdf/25082817.pdf

Stark, B.P. & D.H. Ray. 1983. A revision of the genus

Helopicus (Plecoptera: Perlodidae). Freshwater

Invertebrate Biology, 2(1):16–27.

Stark, B.P. & J.W. Kyzar. 2001. Systematics of

Nearctic Paraleuctra with description of a new

genus (Plecoptera: Leuctridae). Tijdschrift voor

Entomologie, 144(1):119–135.

Stark, B.P. & R.W. Baumann. 2004. The winter

stonefly genus Paracapnia (Plecoptera:

Capniidae). Monographs of the Western North

American Naturalist, 2(1):96–108.

Stark, B.P., S.W. Szczytko & R.W. Baumann. 1986.

North American stoneflies (Plecoptera):

systematics, distribution, and taxonomic

references. Great Basin Naturalist, 46:383–397.

https://www.jstor.org/stable/pdf/41712245.pdf

Stark, B.P., S.W. Szczytko & B.C. Kondratieff. 1988.

The Cultus decisus complex of eastern North

America (Plecoptera: Perlodidae). Proceedings

of the Entomological Society of Washington,

90(1):91–96. http://www.biodiversitylibrary.org/

page/16144505

Stewart, K.W. & B.P. Stark. 1988. Nymphs of North

American stonefly genera (Plecoptera).

Entomological Society of America. Thomas Say

Foundations, Entomological Society of America

Monographs, 12. 460 pp.

Stewart, K.W. & B.P. Stark. 2002. Nymphs of North

American stonefly genera (Plecoptera).

University of North Texas Press (reprint). 400 pp.

Stewart, K.W. & B.P. Stark. 2002. Nymphs of North

American stonefly genera (Plecoptera). Second

Edition. The Caddis Press. Columbia, Ohio. xii +

510 p.

Surdick, R.F. 1985. Nearctic genera of

Chloroperlinae (Plecoptera: Chloroperlidae).

Illinois Biological Monographs 54. University of

Illinois Press, Urbana, Illinois. 146 pp.

Surdick, R.F. 2004. Chloroperlidae (The Sallflies).

Pp. 1–60. In: Stoneflies (Plecoptera) of eastern

North America, Volume II, Chloroperlidae,

Perlidae, and Perlodidae (Perlodinae). B.P. Stark,

B.J. Armitage (Eds.) Bulletin of the Ohio

Biological Survey, Columbus, Ohio. 192 pp.

Szczytko, S.W. & B.C. Kondratieff. 2015. A review of

the eastern Nearctic Isoperlinae (Plecoptera:

Perlodidae) with the description of twenty-two

new species. Monographs of Illiesia, Number

1:1–289. http://illiesia.speciesfile.org/papers/

Monographiae-of-Illiesia.pdf

Page 19: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 48

Verdone, C.J. & B. Kondratieff. 2016. A new species

Isoperla Banks (Plecoptera: Perlodidae) from the

Appalachian Mountains, Virginia & West

Virginia, U.S.A. Illiesia, 12(13):74-85.

http://illiesia.speciesfile.org/papers/Illiesia12-

13.pdf

Unzicker, J.D. & V.H. McCaskill. 1982. Plecoptera.

Pp. 5.1–5.50. In: Aquatic insects and oligochaetes

of North and South Carolina. A.R. Brigham,

W.U. Brigham & A. Gnilka (Eds.) Midwest

Aquatic Enterprises, Mahomet, Illinois.

Woodward, S.L. & R.L. Hoffman. 1991. The nature

of Virginia. Pp. 23–48. In: Virginia’s endangered

species: Proceedings of a symposium. K.

Terwilliger (coordinator). McDonald &

Woodward Publishing Company. Blacksburg,

Virginia. 672 pp.

Appendix A. List of sampling locations from Georgia 2016-2017.

Physiographic

Province County Habitat Location Latitude Longitude

Appalachian Plateau Dade Bear Creek SR 189, Cloudland State Park 34.82823 -85.45934

Appalachian Plateau Dade Daniel Creek Cloudland State Park 34.82457 -85.49074

Appalachian Plateau Walker West Chickamauga Creek SR 136 34.80250 -85.34884

Blue Ridge Dawson Little Amicalola Creek Amicalola Falls State Park 34.57233 -84.24107

Blue Ridge Fannin Noontootla Creek 10.4 km NNE Amicalola Mtn. at FS Rd. 58 34.68791 -84.20182

Blue Ridge Fannin Noontootla Creek 8.9 km NNE Amicalola Mtn. 34.67289 -84.19695

Blue Ridge Fannin Noontootla Creek 9.0 km NNE Amicalola Mtn. 34.67347 -84.19692

Blue Ridge Fannin Seep along FS 58 10.4 km NNE Amicalola Mtn. 34.68595 -84.19587

Blue Ridge Fannin Toccoa River 16.0 km SE Blue Ridge at GA-5 34.75758 -84.20606

Blue Ridge Gilmer Rock Creek 12.6 km NE Ellijay at US-76 34.77922 -84.39091

Blue Ridge Gilmer Rock Creek Rock Creek Rd. 34.78176 -84.33619

Blue Ridge Lumpkin Frogtown Creek Hwy 19, Desoto Falls Rec. Area 34.70649 -83.91634

Blue Ridge Lumpkin Nimblewill Creek Nimblewill Gap Rd. 34.56536 -84.14637

Blue Ridge Murray Conasauga River Old Hwy 2 34.97461 -84.64487

Blue Ridge Murray Holly Creek Camp Rd. Day Use Area 34.81181 -84.66080

Blue Ridge Murray Jigger Creek East Cowpen Rd. 34.96260 -84.63861

Blue Ridge Rabun Chattooga River Hwy 76 Fishing Access 34.81404 -83.30647

Blue Ridge Rabun Greasy Creek North Germany Rd. 34.91460 -83.41972

Blue Ridge Rabun North Fork Chattooga River Hwy 28 34.92010 -83.16924

Blue Ridge Towns headwaters of Soapstone Creek 1.5 km SSE Brasstown Bald 34.85994 -83.80254

Blue Ridge Towns Soapstone Creek 2.9 km SE Brasstown Bald 34.85588 -83.78517

Blue Ridge Union Wolf Creek Vogel S.P. at Burnette Branch 34.76311 -83.92679

Blue Ridge Union Left Fork Nottely River 8.1 km NNW Pinnacle Mountain 34.75055 -83.84511

Blue Ridge Union Nottely River 8.0 km NNW Pinnacle Mountain 34.74962 -83.84646

Blue Ridge White Tributary to Dukes Creek 11.4 km N Cleveland 34.69804 -83.78149

Coastal Plain Bleckley Ocmulgee River James Dykes Park 32.41439 -83.48319

Coastal Plain Houston Burnham Brand Creek Hwy 26, SE of Elko 32.31585 -83.66442

Page 20: STUDIES ON THE STONEFLIES OF GEORGIA WITH …illiesia.speciesfile.org/papers/Illiesia13-03.pdfLeuctridae Leuctra biloba Claassen, 1923 Holotype Leuctridae Leuctra moha Ricker, 1952

Verdone, C.J., B. Kondratieff, R.E. DeWalt, and E. South. 2017. Studies on the stoneflies of Georgia with the description of a new species

of Soyedina Ricker, new state records and an annotated checklist. Illiesia, 13(03):30-49. https://doi.org/10.25031/2017/13.03

Illiesia – http://illiesia.speciesfile.org Volume 13 – Number 3 – Page 49

Coastal Plain Macon Flint River Montezuma Bluffs Park 32.33682 -84.03118

Coastal Plain Stewart Hannahatchee Creek Co Rd. 150 32.14157 -84.75333

Piedmont Plateau Greene Fishing Creek Hwy 15 33.66975 -83.26152

Piedmont Plateau Greene Harris Creek Nichols Rd. 33.72509 -83.23737

Piedmont Plateau Greene Oconee River Hwy 15, Oconee NF campground 33.72149 -83.29095

Piedmont Plateau Habersham trib. to Lake Russell Lake Russell Rd. 34.50077 -83.48914

Piedmont Plateau Paulding Bluffy Creek Hulsey Town Rd. 33.89260 -84.92486

Piedmont Plateau Paulding trib. to Pumpkinvine Creek Hulsey Town Rd. 33.90456 -84.91684

Piedmont Plateau Pickens Talking Rock Creek 6.3 km WNW Talking Rock at GA-136 34.52624 -84.57093

Piedmont Plateau Putnam Big Indian Creek Big Indian Creek Rd. 33.39687 -83.47433

Piedmont Plateau Putnam Little River Glades Rd. 33.37248 -83.47730

Piedmont Plateau Putnam Murder Creek Hillsboro Rd. 33.26861 -83.49739

Piedmont Plateau Putnam Rock Creek Rock Creek Rd. 34.78176 -84.33619

Piedmont Plateau Upson Flint River Sprewell Bluff Park 32.85381 -84.48034

Ridge and Valley Floyd Johns Creek Everett Springs Rd. S of Camp Sydney Drew 34.52889 -85.11405

Ridge and Valley Floyd Johns Creek Everett Springs Rd. 34.56416 -85.10104

Ridge and Valley Floyd trib. to Johns Creek FR 103, The Pocket Rec. Area 34.58443 -85.08587

Ridge and Valley Murray Head Branch Old Hwy 2, 2 mi. E of Cisco 34.94477 -84.71942

Ridge and Valley Murray trib. to Conasauga River East Cowpen Rd. 34.95390 -84.67750

Ridge and Valley Walker Snake Creek SR 138, S. of Snake Creek Gap 34.64711 -85.06106

Ridge and Valley Walker trib. to Johns Creek Keown Falls Trail 34.61380 -85.08757

Submitted 19 July 2017, Accepted 10 August 2017, Published 28

August 2017

Hosted and published at the University of Illinois, Illinois

Natural History Survey, Champaign, Illinois, U.S.A.